North American Fungi

Volume 5, Number 5, Pages 9-21 Published December 22, 2010

Amanita in the Rocky Mountain alpine zone, USA: New records for A. nivalis and A. groenlandica

Cathy L. Cripps1 and Egon Horak2

1Plant Sciences and Plant Pathology Department, Montana State University, Bozeman, MT 59717-3150, U.S.A; 2Nikodemweg 5, AT-6020 Innsbruck, Austria

Cripps, C. L., and E. Horak. 2010. Amanita in the Rocky Mountain alpine zone, USA:New records for A. nivalis and A. groenlandica. North American Fungi 5(4): 9-21. doi:10.2509/naf2010.005.0052

Corresponding author: Cathy L. Cripps, [email protected]. Accepted for publication June 20, 2010. http://pnwfungi.org Copyright © 2010 Pacific Northwest Fungi Project. All rights reserved.

Abstract: A limited number of Amanita species have been reported from cold dominated arctic-alpine environments, primarily with dwarf and shrub willows. This includes reports from Alaska, Canada, Greenland, Iceland, Scandinavia, Scotland, the Alps, and Russia. Here we report Amanita nivalis, A. groenlandica f. alpina n. f. and A. vaginata above tree line in the Rocky Mountains with Salix reticulata, S. nivalis, S. glauca, and S. planifolia. The distribution of Amanita groenlandica is extended to the north-central Rockies, and more definitively to the Beartooth Plateau at 3,100 to 3,400 m a.s.l. A new alpine form is described for this arctic species. Amanita nivalis is confirmed from numerous cirques and high passes 3,600-3,900 m a.s.l. in the southern Rocky Mountains (Front Range, Sawatch Range, San Juan Mountains), adding a disjunct component to its circumpolar distribution. These localities are thousands of miles from the Arctic and are likely near the southern-most extent of its distribution in North America. Rocky Mountain species are reported in context with arctic-alpine Amanita species from the Northern Hemisphere.

Key words: Agaricales, Amanitaceae, Amanita groenlandica, Amanita nivalis, alpine, Arctic, cold- climate fungi, ectomycorrhizal, Rocky Mountains, Salix, tree line 10 Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21

Introduction: The genus Amanita is known from the krummholz conifer zone. All are from arctic-alpine habitats throughout the ectomycorrhizal species associated with alpine Northern Hemisphere (Favre 1955, Gardes and vegetation including shrub willows Salix glauca Dahlberg 1996, Gulden et al. 1985, Hallgrimsson L. and S. planifolia Pursh., and dwarf willows S. 1979, Hutchison et al. 1988, Knudsen & Borgen reticulata L. and S. arctica Pall. Three species of 1987, Miller et al. 1982, Watling 1985, 1987), Amanita are reported here from the Rocky however, there have been no formal reports of Mountain alpine zone along with their ecology strictly alpine species from the Rocky Mountains. and in context with other arctic-alpine taxa in Only a limited number of Amanita species occur this genus. in cold dominated environments mainly in association with dwarf/shrub willows and dwarf birch. At least one species (A. nivalis) has a Materials and Methods northern circumpolar distribution pattern. Study Area Historical taxonomic confusion and unresolved The Rocky Mountains extend along the spine of synonymies have made it unclear as to the exact North America continuing 5,000 km north from number of distinct arctic and alpine taxa of New Mexico through Canada to Alaska. The Amanita. To add to the confusion, the Rocky Mountain alpine tundra, although basidiomes of many alpine Amanita species are fragmented on mountain tops, constitutes a white, near white, or possess a whitish pileus phytogeographically significant region, under some environmental conditions. Amanitas comprised of the Northern, Middle, and Southern often occur as single basidiomes in alpine floristic provinces across a range of latitudes habitats so intraspecific variation cannot be (Arno and Hammerly 1984). Our study examined easily examined. Harsh arctic and alpine sites south of the Canadian border, above treeline conditions can quickly destroy salient features, at 3,000 m a.s.l. in the north at latitude 45°N which is problematic in macromorphological (Montana and Wyoming) and above treeline at evaluation. A majority of arctic-alpine Amanita 3,700 m a.s.l. in southern Colorado at latitudes of species are in sect. Vaginata characterized by 37°-39°N. Portions of the alpine areas in both of basidiomes with a saccate volva, absence of an these regions escaped glaciation. annulus and non-amyloid spores (Tulloss 1994). Species within this group often lack distinct Middle-Northern Floristic zone: The Beartooth features that allow clear taxonomic separation, Plateau is our primary study area for this floristic although molecular work has helped define zone, and extends from Montana into northern subalpine species (Drehmel et al. 1999, Moncalvo Wyoming. The plateau is one of 11 situated along et al. 2000, Weiss et al. 1998). the MT-WY border that together comprise the most extensive alpine tundra in the lower 48 An extensive NSF-sponsored survey of alpine states (Anderson 1994). The Beartooth Plateau fungi along the Rocky Mountain cordillera for the itself is about 20 km long by 10 km wide, at lat. Beartooth Plateau on the Montana/Wyoming 450N and long 109°W. The vast sweeping gentle border and the southern floristic zone in slopes are covered by an intricate patchwork of Colorado resulted in over 2,000 collections of meadows, shrubby wetlands, grasslands and fell- alpine agarics that fall into ca. 200 species fields (Cooper et al. 1997, Eversman 1995, designations (Cripps and Horak 2006a, 2006b, Johnson and Billings 1962). Details of the 2008, Osmundson et al. 2005). Amanita species vegetation and soils are in Anderson (1994), were regularly collected as part of this survey Bamberg and Major (1968), and Scott (1995). from habitats above treeline and strictly away Soils are primarily derived from granitic material. Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 11

The collecting season for fungi is typically from drawn, and identified on subsequent completion mid-July to the end of August. of microscopic examination. Spores and other microscopic features were examined in Southern Floristic Zone: The southern Rocky 3% KOH after rehydration. Colors are from Mountains are isolated from the northern ranges Kornerup and Wanscher (1967). Voucher by the Wyoming Plateau. The individual study specimens are in herbaria at Montana State sites lie in a chain along the Continental Divide at University, Bozeman, MT (MONT) and the elevations from 3600 to 4000 m a.s.l. at lat. 36°- Institute of Integrative Biology, ETH, Zurich, 38°N. The sites include high mountain passes, Switzerland (ZT). An advantage for our study was unique cirques, and basins with a wealth of that particular environmental conditions (high habitats including open windswept slopes, precipitation at local fungus peak season) often shrubby wetlands, extensive dwarf Salix and resulted in large fruitings that provided an Dryas patches, meadows, turf, and fell-fields. A opportunity to examine basidiome variation variety of soil types occur in the mountain ranges within the main alpine taxa that occur in the from granitic to limestone. The fruiting season is Rockies. from mid-July through August or into September after which snow can close the high areas. Results: Numerous basidiomes of Amanita were observed from 1999 and 2008 on the Ectomycorrhizal Vascular Plants: The primary Beartooth Plateau sites (MT/WY) above 3,000 m ectomycorrhizal plants of these alpine areas a.s.l. (Fig. 1) and on the southern sites (Colorado) include dwarf willow species S. reticulata, S. above 3,700 m a.s.l. The records represent three arctica and more rarely S. rotundifolia Trautv. species of Amanita that occur above tree line (not and S. cascadensis Cockll. (Cripps and Eddington in association with conifers) in the Rocky 2005, Scott 1995). Shrub willows consist Mountains. Only North American collections are primarily of S. glauca which is more common in described here with emphasis on local ecology southern areas and S. planifolia which is more and distribution patterns (Table 1). prevalent in the north. Betula glandulosa Michx. (= B. nana L.) is rare in the Rocky Mountain alpine zone and no Amanita species were recorded with this host. Dryas octopetala L. mats can be extensive in the Rocky Mountain alpine belt and host a limited diversity of ectomycorrhizal fungi (Cripps and Horak 2006b), however, to date no Amanita species have been recorded in strict association with Dryas for our sites.

Collection and processing Basidiomes were collected from 1999 to 2008, primarily on the Beartooth Plateau in the North- Central Rockies and on the Front Range, Sawatch Range and San Juan Mountains to the South. Each collection was photographed, described, Fig. 1. Alpine area of Beartooth Plateau, Rocky Mountains, MT/WY, USA. 12 Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21

Table 1. Collection data for Amanita species in the Rocky Mountain alpine zone: A. groenlandica collections are from the Beartooth Plateau* near the Montana and Wyoming border, A. nivalis collections from Colorado**, and A. vaginata from the Beartooths. Cripps (=CLC) collections are in MONT and Horak (=ZT) collections are in ETHZ. Collection Date Range Location Elev. Hosts Amanita groenlandica f. alpina Horak 6410 07 Aug-98 Beartooth Site 1: Birch site 3,100 Salix reticulata Cripps 1213 08 Aug-98 Beartooth Site 2: Highline trail 3,100 Salix spp. Cripps 1377 19 Aug-99 Beartooth Site 1: Birch site 3,000 S. reticulata & S. planifolia Cripps 1392 20 Aug-99 Beartooth Site 2: Highline trail 3,100 Salix reticulata Cripps 1405 21 Aug-99 Beartooth Site 3: Frozen lakes 3,100 Salix glauca/planifolia Cripps 1413 18 July-00 Beartooth Site 2: Highline trail 3,100 Salix reticulata Cripps 1585 20 July-01 Beartooth Site 2: Highline trail 3,100 Salix reticulata Cripps 1779 21 Aug-02 Beartooth Site 4: Headwall 3,400 Salix reticulata Cripps 1929 28 Aug-02 Beartooth Site 2: Highline trail 3,100 Salix cf planifolia Cripps 1970 12 Aug-03 Beartooth Site 5: Top of Pass 3,400 Salix reticulata & S. arctica Cripps 2313 19 July-08 Beartooth Site 2: Highline trail 3,100 Salix spp. Cripps 2328 14 Aug-07 Beartooth Hell-roaring Plateau 3,500 Salix reticulata Amanita nivalis Cripps 1235 12 Aug-98 Front Loveland Pass 3,650 Salix spp. Cripps 1415 08 July-00 San Juan Cinnamon Pass 3,850 Dwarf Salix species Cripps 1640 04 Aug-01 Sawatch Cottonwood Pass 3,700 Salix glauca Cripps 1647 04 Aug-01 Sawatch Cumberland Pass 3,800 Salix glauca Cripps 1665 07 Aug-01 San Juan Mineral Basin 3,650 Salix reticulata (Horak 9763) Cripps 1691 09 Aug-01 San Juan Stony Pass 3,700 S. reticulata & S. arctica (Horak 9534) Cripps 1720 11 Aug-01 San Juan Black Bear Pass 3,900 Salix reticulata Cripps 1727 13 Aug-01 Sawatch Independence Pass 3,650 Not noted Cripps 1773 15 Aug-01 Sawatch Independence Pass 3,650 Not noted Amanita vaginata Horak 6411 07 Aug-98 Beartooth Site 2: Highline trail 3,100 Salix spp.

* Beartooth Plateau sites at 450N, 1090W. **Colorado sites at 37-390N, 1070W. Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 13

Taxonomic descriptions floccose over all, almost pubescent, then breaking up into adder-like pattern (girdles) with a pale Amanita groenlandica Bas ex Knudsen & orange or gray silvery hue; partial veil absent; Borgen f. alpina n.f. (Table 1, Fig. 2) universal veil (volva) saccate, tight around base and then flaring outwards, sometimes in a star Diagnosis. -A typo differt pileo pallide aurantiaco, velo pattern or more ragged, whitish, staining ocher cremeo et odore frucifero. Holotypus varietatis CCL 1377 (MONT). orange in areas or more silvery in age, fragile, easily left in ground, thin or somewhat thick; Amanita groenlandica Bas ex Kundsen & Borgen. AA context white, turning gray in old specimens, Mycology II: 1987: 41-243. stipe hollow. ODOR faintly fruity, becoming = A. groenlandica nom. Prov. Bas in Clèmoncon, 1977:87. unpleasant in age. Illustrations: Knudsen and Borgen 1987: 241; Borgen 1993: 63. BASIDIOSPORES white in deposit, (9.5-)10-12 (-13) x (9-)10-12(-13) μm, with a length range of LR

= 9-13 μm, width range of WR = 9-12 μm, on PILEUS 40-80 (-120) mm wide, hemispherical at average 11.2 x 10.4 μm, QR = 1.0-1.2, Qav = first, then broadly convex, some slightly conic but 1.10, n= 25, globose (or subglobose), hyaline, mostly rounded, margin striate to 10 mm, color smooth, thin-walled, inamyloid, with variable, often white when young, ranging from prominent conic- truncate apiculus, contents pure to mottled combination of cream with often guttulate with one large oil drop. salmon tints, pinkish buff, salmon buff, pale orange (6A4), pale orange brown, pale brown, in BASIDIA 60-80 x 16-20 μm, thin-walled, clavate, age with silver to metallic pale brown 4-spored, sterigmata 3-5 μm long, lacking clamp sheen, more white in dry environmental connections. CHEILOCYSTIDIA occasional, 10-25 conditions, smooth, greasy to kidskin dry; x 22-30 μm, globose, pyriform to clavate, often remnants of universal veil as one piece or as a few with small cell below basal septum, thin-walled, large unevenly scattered patches, dingy cream or lacking clamp connections. buff color, with or without tints of pale orange to pale grayish tan (7C2, 7C3), thin to thick, edges PILEIPELLIS 20-40 μm thick, colorless with sometimes uplifted; margin slightly sulcate in gelatinized suprapellis containing cylindrical some; context thick in pileus, white. hyphae, 3-4 μm wide; subpellis pale yellow brown in KOH, walls semi-gelatinized, interwoven, of LAMELLAE 120-160, free or just touching stipe, filamentous hyphae, 4-6 μm wide, vascular with few lamellulae or none, rarely dichotomous hyphae pale yellow, occasionally dark yellow near cap edge, crowded, broad to 10 mm, brown. PILEUS TRAMA primarily of swollen cells rounded towards margin, white or with faint pale (almost cellular), (10-)20-30 μm wide to 65 μm orange tint, more dingy cream in age; edges finely long, narrowed at the septa, thin-walled, clamp fimbriate, concolorous or darkening to gray in connections absent; vascular hyphae copious, age. pale yellow to dark yellow brown in KOH, 4-10 μm wide, undulating. STIPE 40-100 mm long, 10-25 mm thick, gradually enlarged towards base (slightly HYMENOPHORAL TRAMA of swollen cells, 10- clavate), long, slender, straight to slightly curved, 20(-40) µm wide with narrowed septa, thin- buried in soil or above ground, at first white with walled. SUBHYMENIUM of short squarish cells in or without pale orange cream tints, in age with chains 10 x 10 μm. UNIVERSAL VEIL primarily of more gray or silvery tones, surface at first filamentous cylindrical hyphae, 4-6 μm wide, 14 Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 thin-walled, with y-branching, interwoven, OBSERVATIONS: Taxonomically Amanita septate, intermixed with inflated cells groenlandica is in sect. Vaginatae based on (sphaerocysts) 15-20 x 32-65 μm, in localized inamyloid spores and lack of a partial veil. The areas up to 75% of veil tissue, mostly in outer Rocky Mountain alpine form (forme alpina) is tissue, clamp connections absent. robust with coloration reminiscent of a “cream- sicle” that varies from solid to mottled color HABITAT AND DISTRIBUTION: Scattered in mats combinations of cream, pale orange, and pale of dwarf willow Salix reticulata and near shrub silvery brown. It is similar to the arctic form of willows S. planifolia and S. glauca in the alpine Amanita groenlandica Bas ex Knudsen & T. vegetation zone. It fruits from July through Borgen (1987) personally observed in several August, above tree line at elevations of 3,100- localities in Greenland. The Rocky Mountain 3,400 m a.s.l. The alpine form occurs prolifically alpine form differs from the arctic form seen in on the Beartooth Plateau near the Wyoming/ Greenland by its taller stature, different Montana border and was not recorded from the coloration, and less viscid pileus. The alpine form southern Rockies. is more in the pale orange brown range and is not typically dominated by gray brown or snuff SPECIMENS EXAMINED: USA. MONTANA, Carbon brown colors as described for the arctic form County: Beartooth Plateau, 8 July 1998, E. Horak (Knudsen and Borgen 1987). The universal veil is 6410 (ZT); 19 Aug 1999, C.L. Cripps 1377 (FORM typically more cream color (not as gray as the TYPE, MONT, MycoBank Number 518728); 14 arctic form), but can be in the gray range. July 2001, C.L. Cripps 1568 (MONT); 28 Aug Comparison of dried specimens shows a 2002, C.L. Cripps 1929 (MONT); Hell-roaring difference on reconstitution in KOH, with the Plateau, 14 Aug 2007, C.L. Cripps 2328 (MONT). arctic form veil exhibiting darker gray coloration. WYOMING, Park County: Beartooth Plateau, 8 The odor of the alpine form is definitely fruity Aug 1998, C.L. Cripps 1213 (MONT); 20 Aug 1999, when young then becomes unpleasant in age, C.L. Cripps 1392 (MONT); 21 Aug 1999, C.L. whereas specimens from the arctic are described Cripps 1405 (MONT); 18 July 2000, C.L. Cripps as odorless. For our sites, this species only has 1413 (MONT); 20 July 2001, C.L. Cripps 1585 been observed on the Beartooth and Hell-roaring (MONT); 21 Aug 2002, C.L. Cripps 1779 (MONT); Plateaus where it fruits in large numbers as a 12 Aug 2003, C.L. Cripps 1970 (MONT). conspicuous element of the alpine mycota.

Fig. 2. Amanita groenlandica Bas ex Knudsen forme alpina (CLC 2328) in the Rocky Mountains. Half life size. Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 15

Amanita nivalis Grev. Scot. Crypt. Fl. 1(4): pl. exterior can stain slightly ochre. Context white. 18. 1821. (Table 1, Fig. 3). Odor not distinctive.

Amanitopsis nivalis (Grev.) Sacc. Syll. Fung. 5: 22. 1887. BASIDIOSPORES white in deposit, 10-12(-14) x

Amanitina nivalis (Grev.) E.-J. Gilbert. Icon. Mycol. 27 8.5-11 μm, with a length range of LR=9-14 μm and

(suppl. 1): 78. 1941. width range of LR= (7.5)8.5-13.5 μm, on average

Amanita vaginata var. nivalis (Grev.) E.-J.Gilbert. Le genre 10.7 x 9.7 μm, Qav = 1.1, QR = 1-1.2(1.4), n = Amanita Persoon (Lons le-Saunier): 141. 1918. 25, globose or subglobose, colorless, smooth, Amanitopsis vaginata var. nivalis (Grev.) Peck. Ann. Rep. thin- walled, inamyloid, with prominent N.Y.State Mus. 47: 169. 1884. apiculus; content often guttulate with one Amanita vaginata f. nivalis (Grev.) Vesely. Ann. Mycol. 31: large oil drop. 279. 1933. Amanita vaginata f. oreina J. Favre. Ergebn. wissensch. BASIDIA: 70-80 x 15-18(20) μm, clavate, thin- Unters. Schweiz. NatParks (5) 33: 205. 1955. walled, 4-spored, sterigmata 3-6 μm long, clamp connection absent at base. STERILE ELEMENTS Illustrations: Gulden et al. (1985: 25), Moser and Jülich (paracystidia) on lamellar edges globose or (1988: 1, 2), Campo and Bizio (2000: 177, 179, 182), Tulloss clavate, 15-30 x 10-20 μm, colorless. (2005), Jamoni 2008: 166-168. PILEIPELLIS 20-30 μm in depth, hyaline, with gelatinized colorless suprapellis containing PILEUS: 30-70 mm wide, convex to hyphae 3-4 μm wide. Subpellis pale yellow brown hemispherical when young, becoming shallow in KOH, semi-gelatinized, thick, individual convex, to almost applanate, mostly without hyphae 5-17 μm in diameter, cylindrical, distinct umbo, margin distinctly striate or almost branched, without clamp connections; vascular plicate-striate; color variable, ranging from hyphae 5-7 μm wide, with yellow contents in whitish silver, pale gray, pale silver to pale gray KOH, undulating, occasionally branching. brown (4C2), smooth, kidskin to slightly greasy. HYMENOPHORAL TRAMA bilateral, of inflated Universal veil typically absent on pileus surface, hyphae 12-18 μm wide with constricted septa and occasionally as one patch, in a few collections as occasional vascular hyphae same as for scattered warts (pyramidal or not) possibly in dry pileipellus. SUBHYMENIUM: of short cells, some conditions, variable within a local population or inflated. UNIVERSAL VEIL composed mostly of among collections, tissue whitish, staining pale filamentous hyphae, 4-6 μm wide and up to 40 gray. Context thin in pileus, white. μm long, a few scattered pyriform sphaerocysts 20-40 x 30-40 μm present, clamp connections LAMELLAE: L=100, free or just touching stipe, absent. typically without lamellulae; crowded, white, some with a pink cast especially when young but HABITAT AND DISTRIBUTION: All collections are fading in age, drying pale orange, edges floccose, from the southern Rocky Mountains (CO) and it concolorous or graying. was found at over half of the localities visited in that state (Table 1). No representatives are STIPE: 40-60(-80) mm long, 5-10(-13 at base) reported here for northern sites however a recent mm thick, gradually enlarged towards base, collection from the Beartooth may represent this hollow, white or with pale gray cast, minutely taxon. Amanita nivalis is a well known arctic- floccose for whole length with longitudinal pruina alpine species associated with willows; it has a (not adder-like pattern). Partial veil absent. circumpolar distribution and is now officially Universal veil saccate, flaring, lobed, white, confirmed from the Rocky Mountains and lower 48 states. 16 Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21

SPECIMENS EXAMINED: USA. COLORADO: absent to a few patches, and in a few collections Summit County, Front Range, Loveland Pass, 12 scattered over the whole pileus as warts (also Aug 1998, C.L. Cripps 1235 (MONT); San Juan described by Knudsen and Borgen 1987). The County, San Juan Mountains: Cinnamon Pass, 8 fruiting body is less robust than A. groenlandica July 2000, C.L. Cripps 1415 (MONT); Mineral and the pileus more striate. Lamellae can be Basin, 7 Aug 2001, E. Horak 9763 (ZT, MONT bright pink when young, changing to pale orange, 1665); Stony Pass, 9 Aug 2001, C.L. Cripps 1691 and darkening on drying. The stipe is pruinose (MONT); Black Bear Pass, 11 Aug 2001, C.L. and lacks the adder-like pattern of A. Cripps 1720 (MONT); Chaffee/Gunnison groenlandica. County, Sawatch Range: Cottonwood Pass, 4 Aug 2001, C.L. Cripps 1640 (MONT); Amanita cf. vaginata (Bull.) Lam. Cumberland Pass, 4 Encycl.Méth. Bot. (Paris) 1: 109. 1783 (Table 1). Aug 2001, C.L. Cripps 1647 (MONT); Pitkin/Lake County, Independence Pass, 13 Aug 2001, C.L. PILEUS 60 mm in diameter, convex to broadly Cripps 1727 (MONT); same locality, 15 Aug 2001, umbonate-expanded, uniformly lead grey, ash C.L. Cripps 1773 (MONT), E. Horak 9534 (ZT). grey to fuliginous, margin crenate-splitting, if moist subviscid, mat in dry condition, with OBSERVATIONS: Amanita nivalis is a small whitish lumps from veil. LAMELLAE free, dense, delicate species restricted to arctic and alpine lamellulae scattered, white, floccose pale grey habitats with willows. North American species edges. STIPE -80 x -18 mm, gradually exhibit the whole range of color variations from attenuated towards apex, uniformly grey, whitish to silvery gray or pale gray brown. There conspicuously longitudinal striate, not adder are wide swings in temperature and humidity in patterned, minutely powdery-pruinose, hollow, the alpine compared to the arctic, and remnants solitary. of the universal veil on the cap can vary from ANNULUS absent. VOLVA saccate, persisting,

Fig. 3. Amanita nivalis Grev. (CLC 1691=EH 9534), darker form, ZT 9808 is paler form: specimens from Colorado. Strip = 2 cm. Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 17 white (with very pale tinge). CONTEXT white. in arctic-alpine areas of Greenland (Watling ODOR not distinctive. SPORES white, 9.5-11 µm, 1977, 1983, Knudsen & Borgen 1987), Iceland (sub-) globose. VOLVA composed of cylindrical (Hallgrimsson 1972, 1979), Scandinavia (Bas hyphae, rarely intermixed with ovoid or globose 1977, Jacobsson 1984, Gulden et al. 1985), the cells. Austrian Alps (Gerold 1986, Peitner et al. 1999), the French Alps (Bon 1985), the Italian Alps SPECIMEN EXAMINED: USA: Montana, (Scaramella 1930, Bon 1987, Jamoni 2008), the Carbon County, Beartooth Plateau, Highline Swiss Alps (Boudier & Fischer 1895, Favre 1955, Trail, alpine habitat, on soil associated with Kúhner 1972, Breitenbach & Kränzlin 1991, Salix spp., 7 Aug. Senn-Irlet 1992), Alaska (Kobayasi 1967), and 1998, E. Horak 6411 (ZT). Russia (Knudsen & Mukhin 1996, Karatygin et al. 2008). OBSERVATIONS: This species is reported once on the Beartooth Plateau with Salix and fits the There is some taxonomic confusion in the description for the typical form of A. vaginata. interpretation of this species and several names This collection is characterized by a lead grey may be synonymous (see Bas 1982, Campo & pileus, white lamellae with pale grey floccose Bizio 2000, Knudsen & Borgen 1987, Tulloss edges, velutinous stipe without adder pattern 1994, Watling 1985, 1987), thus we prefer to and whitish saccate volva. It is also known from treat this taxon in the broad sense here subalpine habitats in the region with aspen. (Knudsen & Borgen 1987). Amanita species with the nomens of oreina and hyperborea, whether Discussion: The present contribution is the as a species, subspecies, or form need further first formal in-depth report of Amanita nivalis taxonomic clarification beyond the scope of this and Amanita groenlandica from the Rocky paper (Bon 1987). There are several similar Mountains south of the Canadian border and a species of white to gray Amanita in the Rocky new form is described for the latter. Mountains, but they occur in subalpine habitats with other host trees (Tulloss 1994). Amanita nivalis is reported from several high- elevation basins and passes in the Front Range, The alpine form of A. groenlandica reported the San Juan Mountains and the Sawatch Range, here from northern sites on the Beartooth all in Colorado from 3,650-3,900 m a.s.l., in Plateau has not been recorded yet from the association with willows, primarily Salix southern Rocky Mountains. It appears reticulata and S. glauca. There is only one morphologically close to Amanita groenlandica possible report of this species from northern which was first described from Greenland sites despite intensive collecting efforts. (Borgen 1993, Knudsen & Borgen 1987). There Colorado may be close to the southern extent of are also reports of A. groenlandica from Canada this arctic-alpine species. (Hutchison et al. 1988; Ohenoja 1972), Norway and Russia (Knudsen and Mukhin 1996) (Fig. 4). Amanita nivalis is the most well-known arctic- The arctic form observed by us in Greenland is alpine species of Amanita and has a circumpolar in our opinion distinct by different colors and distribution with disjunct populations in alpine lack of odor. The alpine form does not extend areas of mountain ranges across the Northern into the subalpine zone with conifers in the Hemisphere (Fig. 4). The “ringless snow Rocky Mountains. Amanita” was originally described from the Scottish Highlands (Greville 1822, reviewed in Other species of Amanita reported from Arctic- Watling 1985, 1987) and subsequently was found alpine habitats include A. arctica Bas, Knudsen 18 Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 and Borgen. This white robust species was first close examination most appear associated with described from Greenland (Knudsen & Borgen conifers in the krummholz zone. These records 1987) and has been recently found in Finland are not considered strict arctic- alpine (PA Moreau, unpublished). The first author inhabitants by our definition. personally observed this collection, and it represents a species not yet recorded for the Acknowledgments: This project was funded Rocky Mountains. However, there is one by the National Science Foundation Biotic unconfirmed report of an alpine Amanita by this Surveys and Inventories program, and we thank name from Eastern Canada (Hutchison et al. them for continuing this program which seeks to 1988). Amanita mortenii Knudsen & Borgen discover and document life on earth. Authors from Greenland appears restricted to Betula thank students Todd Osmundson and Leslie shrubs (Elborne & Knudsen 1990) which are not Eddington for participation in this survey and common in the Rocky Mountains alpine zone. Almut Horak and Don Bachman for their Additional Amanita taxa are occasionally collecting efforts. reported from cold-dominated climates, but on

Amanita groenlandica f. alpina

Amanita nivalis Grev.

Fig. 4. Distribution of Amanita nivalis and Amanita groenlandica in the Northern hemisphere. Cripps & Horak. Amanita, new records, Rocky Mountain alpine. North American Fungi 5(5): 9-21 19

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